Two subtypes of GTPase-activating proteins coordinate tip growth and cell size regulation in Physcomitrium patens.

The establishment of cell polarity is a prerequisite for many developmental processes. However, how it is achieved during tip growth in plants remains elusive. Here, we show that the RHO OF PLANTs (ROPs), ROP GUANINE NUCLEOTIDE EXCHANGE FACTORs (RopGEFs), and ROP GTPASE-ACTIVATING PROTEINs (RopGAPs) assemble into membrane domains in tip-growing cells of the moss Physcomitrium patens. The confinement of membrane domains requires redundant global inactivation of ROPs by PpRopGAPs and the PLECKSTRIN HOMOLOGY (PH) domain-containing RenGAP PpREN. Unexpectedly, PpRopGAPs and PpREN exert opposing effects on domain size and cell width upon overexpression. Biochemical and functional analyses indicate that PpRopGAPs are recruited to the membrane by active ROPs to restrict domain size through clustering, whereas PpREN rapidly inactivates ROPs and inhibits PpRopGAP-induced clustering. We propose that the activity- and clustering-based domain organization by RopGAPs and RenGAPs is a general mechanism for coordinating polarized cell growth and cell size regulation in plants.

ROP GTPase-dependent polarity establishment during tip growth in plants.

Polar cell growth in plants requires a cell peripheral region that undergoes membrane extension and cell wall remodeling. Since the 1990s, RHO-RELATED GTPASES FROM PLANTS (ROPs) have been identified as master regulators that determine the site of cell growth. ROPs function to regulate actin and microtubule cytoskeletons, calcium gradients, and exocytosis, thus directing the delivery of materials for membrane and cell wall extension. In recent years, our understanding of the regulatory mechanisms underlying polar localization and the activation of ROPs has greatly advanced. Evidence points to the crucial roles of membrane lipids, receptor-like kinases, and cell wall components. In this review, we provide updates on the mechanisms underlying polarity control in tip-growing cells, with a focus on ROP effectors and membrane-associated signals. By integrating knowledge from pollen tubes, root hairs, and findings in bryophyte protonema cells and rhizoids, we hope to offer important insights into a common conceptual framework on polarity establishment governed by intercellular and extracellular signals.